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Biodiversity Is Autocatalytic

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Biodiversity Is Autocatalytic Ecological Modelling 346 (2017) 70–76 Contents lists available at ScienceDirect Ecological Modelling j ournal homepage: www.elsevier.com/locate/ecolmodel Biodiversity is autocatalytic a,∗ b c Roberto Cazzolla Gatti , Wim Hordijk , Stuart Kauffman a Biological Diversity and Ecology Laboratory, Bio-Clim-Land Centre of Excellence, Tomsk State University (TSU), Tomsk, Russia b Konrad Lorenz Institute for Evolution and Cognition Research, Klosterneuburg, Austria c Institute for Systems Biology, Seattle, WA, USA a r t i c l e i n f o a b s t r a c t Article history: A central question about biodiversity is how so many species can coexist within the same ecosystem. The Received 14 October 2016 idea that ecological niches are critical for the maintenance of species diversity has received increasing sup- Received in revised form 5 December 2016 port recently. However, a niche is often considered as something static, preconditioned, and unchanging. Accepted 6 December 2016 With the “Biodiversity-related Niches Differentiation Theory” (BNDT), we recently proposed that species themselves are the architects of biodiversity, by proportionally increasing the number of potentially Keywords: available niches in a given ecosystem. Autocatalytic sets Along similar lines, but independently, the idea of viewing an ecosystem of interdependent species as Ecological niches Biodiversity an emergent autocatalytic set (a self-sustaining network of mutually “catalytic” entities) was suggested, where one (group of) species enables the existence of (i.e., creates niches for) other species. Here, we show that biodiversity can indeed be considered a system of autocatalytic sets, and that this view offers a possible answer to the fundamental question of why so many species can coexist in the same ecosystem. In particular, we combine the two theories (BNDT and autocatalytic sets), and provide some simple but formal examples of how this would work. © 2016 Published by Elsevier B.V. 1. Introduction ecological niche is the role and the position a species has in its environment (its food and shelter needs, its survival and reproduc- The variability among living organisms in terrestrial, marine and tion strategies, etc.). The concept of a niche as the set of ecological other aquatic ecosystems, and the ecological complexes of which requirements, from the reproductive to the alimentary ones, devel- they are a part, have been defined with the term “biodiversity” (CBD oped by Elton (1927) and improved by Hutchinson (1957) with the Secretariat, 1992). Apart from the formal definitions and the dif- definition of hyper-volume, is a powerful tool for understanding ferent ways to measure it, the central question about biological the role of each species in its environment. diversity on Earth is how so many species can coexist within the These multidimensional spaces or hypervolumes that include same ecosystem (Sherratt and Wilkinson, 2009). all of a species’ interactions with the biotic and abiotic factors of In an attempt to explain this issue, some authors formalized neu- its environment, led to the consideration of niches as fundamental tral theories of biodiversity (MacArthur and Wilson, 1967; Hubbell, ecological variables able to regulate species composition and rela- 2001), which assume that all species belonging to the same trophic tions within an ecosystem. For example, it has been suggested that level of an ecological community are “neutral” in relation to their niche differences stabilize competitor dynamics by giving species fitness. This implies that there are no real differences between the higher per-capita population growth rates when rare than when niches of each species and that their success is dictated by the common, and that coexistence occurs when these stabilizing effects randomness of the moment (Rosindell et al., 2011). of niche differences overcome species in overall competitive ability In contrast, the idea that niches are critical for the maintenance (Levine and HilleRisLambers, 2009). Moreover, it seems that nest- of species diversity, challenging the neutral theory of biodiver- edness of niches reduces interspecific competition and enhances sity, has received increasing support recently (McGill, 2003). An the number of coexisting species (Bastolla et al., 2009). Some authors suggested a relationship between the utilization of ecospace and change in diversity of, for example, marine shelf faunas through time (Bambach, 1983). However, most of these ∗ Corresponding author. previous studies emphasized the effect of niche partitioning as a E-mail addresses: [email protected] (R.C. Gatti), global long-term pattern in the fossil record to explain the expo- [email protected] (W. Hordijk), [email protected] nential diversification of life (Benton and Emerson, 2007). The main (S. Kauffman). http://dx.doi.org/10.1016/j.ecolmodel.2016.12.003 0304-3800/© 2016 Published by Elsevier B.V. R.C. Gatti et al. / Ecological Modelling 346 (2017) 70–76 71 explanation for a pattern of exponential diversification is that as entire set is able to sustain and reproduce itself from a basic food diversity increases, the world becomes increasingly divided into source. In other words, the set as a whole is self-sustaining and col- finer niche spaces. This explanation could be a result of the fact that lectively autocatalytic. This concept is intimately related to those nearly all studies of the impact of species interactions on diversi- of Ulanowicz (2008) and Maturana and Varela (1980), but worked fication have concentrated on competition and predation, leaving out in more mathematical detail (Hordijk, 2013). out the importance of other types of interactions (Joy, 2013). Autocatalytic sets were originally defined in the context of However, the idea that interactions between species are impor- chemistry (in particular polymer systems; see below), but have tant catalysts of the evolutionary processes that generate the more recently been extended to study systems in biology (Sousa remarkable diversity of life is gaining interest among ecologists. For et al., 2015) and possibly economics (Hordijk, 2013). Here, we show instance, it has been shown that symbiosis between gall-inducing that biodiversity can also be considered a system of autocatalytic insects and fungi catalyzed both the expansion in resource use sets, and that this view offers a possible answer to the fundamental (niche expansion) and diversification (Joy, 2013). Indeed, facilita- question of why so many species can coexist in the same environ- tion (a process that allows the colonization and presence of new ment. species taking advantage of the presence of other ones by expand- In the following sections, we briefly review the Biodiversity- ing the ecosystem hypervolume) plays a major role in species related Niches Differentiation Theory (BNDT) and the theory of coexistence, strongly increasing the biodiversity of an area. With autocatalytic sets. The BNDT describes how the number of species in the “Biodiversity-related Niches Differentiation Theory” (BNDT), an ecosystems changes over time, depending on the number cur- we recently proposed that species themselves are the architects rently present, and autocatalytic sets can provide a mechanistic of biodiversity, by proportionally (possibly even exponentially) explanation for this process. This idea is illustrated with a simple increasing the number of potentially available niches in a given but formal example. ecosystem (Cazzolla Gatti, 2011). Fath (2007) suggested that all objects in ecological networks 2. The biodiversity-related niches differentiation theory interact with and influence the others in the web and that there are no null community-level relations. Moreover, network mutualism With the Biodiversity-related Niches Differentiation Theory is made by community-level relations that usually have a greater (BNDT) (Cazzolla Gatti, 2011), we recently proposed that species occurrence of mutualism than competition, making them more themselves are the architects of the greatest biodiversity of a given positive than the direct relations that produced them. Fath (2014) environment, because through the realization of their fundamen- also proposed that there are no individual species as such, but only tal niche they allow for an expansion of available niches for other historically contingent constructs that emerge from the structural species. The BNDT states that (Cazzolla Gatti, 2011): couplings of physical and environmental systems. Species them- . “ . .in natural conditions of immigration and emigration, with selves, within an ecosystem, appear and disappear over time, as the every environmental condition, species tend – directly or indi- environmental conditions allow and construct. A species emerges rectly, thanks to their simple presence and life roles – to increase from this environment and is an expression, in fact a historically the number of potentially available niches for the colonization of contingent expression, of those interactions. In other words, species other species, enhancing the limit imposed by the basal hyper- are expressed and maintained by a complex interacting ecological volume, until they reach the carrying capacity of the ecosystem. network. At the same time, niches and mutualistic networks of the ecosys- Paraphrasing von Uexküll (1926), the output of one species, tem allow, through circular and feedback mechanisms, the rise through a series of direct linkages, indirectly connects back again of the number of species,
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